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<I>Stigmaphyllon</I> (<I>Malpighiaceae</I>)

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<I>Stigmaphyllon</I> (<I>Malpighiaceae</I>) Blumea 56, 2011: 73 –104 www.ingentaconnect.com/content/nhn/blumea RESEARCH ARTICLE doi:10.3767/000651911X573444 Revision of Ryssopterys and transfer to Stigmaphyllon (Malpighiaceae) C. Anderson1 Key words Abstract Molecular evidence shows the Old World genus Ryssopterys (Malpighiaceae) nested within the New World genus Stigmaphyllon; therefore, Ryssopterys is here transferred to Stigmaphyllon as subg. Ryssopterys. The androdioecy subgenera share most vegetative and fruit characters. Subgenus Stigmaphyllon comprises 92 species character- Australia ized by hermaphrodite, bilaterally symmetrical flowers. Subgenus Ryssopterys includes 21 species that appear dioecy androdioecious but are probably functionally dioecious; the flowers are either hermaphrodite but likely functionally Indonesia female, owing to inaperturate pollen, or male with a rudimentary gynoecium. All species have radially symmetrical Malpighiaceae flowers in which all parts of each floral whorl are equal; they lack calyx glands as well as the stylar folioles common New Caledonia in subg. Stigmaphyllon, for which the genus is named. The range of subg. Ryssopterys encompasses Indonesia New Guinea (except Borneo and Sumatra), New Guinea, Queensland (Australia), New Caledonia, Vanuatu, the Solomon Islands, Ryssopterys Micronesia, Palau, and the Philippines; S. timoriense has also been recorded from Taiwan and the Ryukyu Islands. An Stigmaphyllon overview of the two subgenera is given. For subg. Ryssopterys summaries of the taxonomic history and morphology, as well as descriptions, a subgeneric key and regional keys, distribution maps, and illustrations of the novelties are provided. Twelve new combinations are proposed: Stigmaphyllon subg. Ryssopterys, S. abutilifolium, S. albidum, S. angustifolium, S. australiense, S. dealbatum, S. discolor, S. grandifolium, S. gymnopodum, S. intermedium, S. taomense, S. timoriense. Ten new species are described: S. brassii, S. mackeeanum, S. mariae, S. mcphersonii, S. merrillii, S. micranthum, S. papuanum, S. pullenii, S. solomonense, S. sundaicum. Published on 12 April 2011 INTRODUCTION activity. Future workers may uncover the origins and phylo- geny of subg. Ryssopterys, the developmental changes in the The family Malpighiaceae is of New World origin, and the Old shift in floral architecture, and the role of dioecy in population World representatives are derived from New World ancestors structure. (Cameron et al. 2001, Davis et al. 2001). Recent investiga- tions of the generic phylogeny of the Malpighiaceae by Davis TAXONOMIC HISTORY & W.R.Anderson (2010) confirmed that the Old World genus Ryssopterys is nested within the New World genus Stigmaphyl­ Genera in Malpighiaceae were very broadly interpreted until lon; all other Old World genera are sister to a New World Adrien de Jussieu began his monographic studies of the family clade. Support values for the Stigmaphyllon clade and for the and proposed clearly defined generic limits. The first species Ryssopterys clade are 100 %. This result is not surprising, if now assigned to subg. Ryssopterys were described in Ban­ one considers that Stigmaphyllon and Ryssopterys differ only isteria L. (Ventenat 1808, De Candolle 1824), Heteropterys in floral morphology and geographical distribution. Because Kunth (Blume 1825), and Hiraea Jacq. (Blume 1825). Jus- retaining Ryssopterys as a separate genus would render sieu (1838) erected the genus Ryssopterys with one species, Stigmaphyllon paraphyletic, the transfer of Ryssopterys to R. timoriensis, based on Banisteria timoriensis DC.; he cited Stigmaphyllon as subg. Ryssopterys is proposed. The diverse Blume’s names Hiraea ovata and H. obscura in synonymy. In subg. Stigmaphyllon includes 92 species (C. Anderson 1997, his Malpighaceaerum synopsis (Jussieu 1840), he added one 2000, 2009) and likely will be divided into additional subgenera new species, R. abutilifolia, and proposed one new combina- as the species relationships become known. tion, R. tiliifolia, based on Ventenat’s Banisteria tiliifolia, with Transfer of Ryssopterys to Stigmaphyllon prompted a review Blume’s Heteropterys albida cited in synonymy. of the Old World species, which revealed a surprising number Jussieu’s monograph of the Malpighiaceae was published in of novelties. As noted under Taxonomic History, the encom- 1843, and under Ryssopterys he listed six species. To R. abu­ passing view presented in Flora Malesiana (Jacobs 1955), tilifolia, R. tiliifolia (now excluding Heteropterys albida), and which assigned all collections from the region covered to either R. timoriensis, he added three new species from the Philippines. R. tiliifolia or R. timoriensis, obscured the true diversity. Sub- Ryssopterys cumingiana and R. dealbata are each based on a genus Ryssopterys now comprises 21 species, of which ten Cuming collection; for R. microstema Jussieu listed, in addition are here newly described. This revision, based entirely on to a Cuming collection, a Blume collection from Java and now herbarium material (see Acknowledgements), must be viewed cited Heteropterys albida Blume in synonymy, thus creating a as a preliminary assessment. Others may find some of my superfluous name. During the next sixty years several species circumscriptions also too broadly drawn; field work will define were added. Hasskarl (1858) proposed R. chrysantha, and the species ranges, which now reflect mostly limited collecting Turczaninow (1863) published R. ovata and R. rufescens, but 1 University of Michigan Herbarium, 3600 Varsity Drive, Ann Arbor, MI 48108 both belong to Aspidopterys A.Juss. Hochreutiner (1904) added USA; e-mail: [email protected]. R. intermedia and Gandoger (1913) R. discolor. © 2011 Nationaal Herbarium Nederland You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 74 Blumea – Volume 56 / 1, 2011 Niedenzu, the second monographer of the Malpighiaceae, but the anterior sepal is eglandular (except in S. boliviense published on Old World Malpighiaceae in 1915 and recognized C.E.Anderson and S. coloratum Rusby). The corolla consists of eight species in Ryssopterys: R. abutilifolia (including R. inter­ four lateral petals and a posterior petal, the ‘flag’; all are clawed media as var. intermedia), R. dealbata (including R. cuming­ (Fig. 1e), but the claw of the posterior petal is longer and stouter iana), R. microstema, R. tiliifolia, R. timoriensis, and three novel- than that of lateral petals. The bee grasps the claw of the ‘flag’ ties, R. angustifolia, R. australiensis, and R. austrocaledonica. petal with its mandibles and reaches between the lateral petals In his treatment of Ryssopterys for Das Pflanzenreich (1928), to scrape the oil glands that are borne on each lateral sepal. Niedenzu also accepted eight species but with some differ- Most species of subg. Stigmaphyllon have a heteromorphic ences; R. abutilifolia, R. angustifolia, R. austrocaledonica, R. androecium (Fig. 1f) and styles with apical folioles (Fig. 1g); dealbata, R. timoriensis, and R. tiliifolia (including R. chrysan­ the anterior style differs from the two equal posterior ones. tha) remained unchanged. He was unaware of Baker’s (1921) The stigma is placed at the apex of the style but always on the publication of R. taomensis, but he accepted R. discolor (de- adaxial angle, never terminally. scribed from New Caledonia) and listed his R. australiensis as In contrast, the flowers of subg. Ryssopterys are radially sym- a synonym. He expanded R. discolor to include all collections metrical (Fig. 1b); the components of each floral whorl are equal. from the Philippines with pubescent leaves, except Cuming The sepals are eglandular (very rarely 1 or 2 sepals of a calyx 1845, which he listed as R. albida; Merrill (1923) had published with a rudimentary gland), and the petal claw is rudimentary or the combination R. albida to replace the superfluous name R. absent. The stamens and styles are uniform, and the stigma is microstema. terminal (Fig. 1c, d). It is assumed that this architecture reflects Except for the addition of R. gymnopoda (Guillaumin 1932) the shift in pollination syndrome in the ancestor of subg. Rys­ and R. grandifolia (Guillaumin 1942), Niedenzu’s interpretation sopterys to one in which the reward is pollen, given the absence of the genus remained accepted until the publication of the of oil bees that visit Malpighiaceae in the Old World. treatment of the Malpighiaceae for Flora Malesiana (Jacobs In previous accounts, such as in Flora Malesiana (Jacobs 1955), 1955). For Ryssopterys Jacobs recognized only two species: subg. Ryssopterys was described as androdioecious, because R. tiliifolia, defined by apiculate anthers and a samara 4–6 cm plants bear either perfect or male flowers. A study of the breed- long, and R. timoriensis (“a very variable species”) for all other ing system of subg. Ryssopterys is beyond the scope of this
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